The phylogeny of Syntrichia: An ecologically diverse clade of mosses with an origin in South America.

PREMISE: To address the biodiversity crisis, we need to understand the evolution of all organisms and how they fill geographic and ecological space. Syntrichia is one of the most diverse and dominant genera of mosses, ranging from alpine habitats to desert biocrusts, yet its evolutionary history remains unclear. METHODS: We present a comprehensive phylogenetic analysis of Syntrichia, based on both molecular and morphological data, with most of the named species and closest outgroups represented. In addition, we provide ancestral-state reconstructions of water-related traits and a global biogeographic analysis. RESULTS: We found 10 major well-resolved subclades of Syntrichia that possess geographical or morphological coherence, in some cases representing previously accepted genera. We infer that the extant species diversity of Syntrichia likely originated in South America in the early Eocene (56.5-43.8 million years ago [Mya]), subsequently expanded its distribution to the neotropics, and finally dispersed to the northern hemisphere. There, the clade experienced a recent diversification (15-12 Mya) into a broad set of ecological niches (e.g., the S. caninervis and S. ruralis complexes). The transition from terricolous to either saxicolous or epiphytic habitats occurred more than once and was associated with changes in water-related traits. CONCLUSIONS: Our study provides a framework for understanding the evolutionary history of Syntrichia through the combination of morphological and molecular characters, revealing that migration events that shaped the current distribution of the clade have implications for morphological character evolution in relation to niche diversity.

Homogenization of bryophyte species after alpine grassland restoration.

The causes of decreasing plant species richness include abandonment of traditional management and the spread of invasive species, even in alpine habitats. Studies on the restoration and management of alpine habitats are predominantly focused on vascular plants, although an important part of alpine vegetation and its diversity is formed by bryophytes. We used bryophytes to indicate changes that occur after the clearcutting of nonindigenous dwarf pine (Pinus mugo Turra) and attempted to reveal the community to which the development of bryophyte species structure was directed. We compared species richness and composition between surveys to test for changes in spatial heterogeneity bryophyte communities. We also tried to reveal the main ecological drivers of the restoration process. The study was performed in the (sub)alpine area of the Eastern High Sudetes Mts. (the Czech Republic). We estimated bryophyte species cover and compared the composition of the bryophyte community in autochthonous grassland areas, areas under the dwarf pine canopy, and clearcut areas to reveal the pattern of shifts 9 years after the treatment. We also measured soil characteristics to reveal the environmental habitat conditions. Evidence of taxonomic homogenization of habitat after dwarf pine removal was found. Light conditions and attributes of litter were the driving factors of successional changes in the bryophyte communities, which led to taxonomic homogenization. This finding explains the slow restoration process due to dwarf pine legacy on the clearcut area. The succession trends were also shaped by unobserved factors, such as climate change and environmental eutrophication. We highly recommended active management and long-term monitoring.

Liverwort oil bodies: diversity, biochemistry, and molecular cell biology of the earliest secretory structure of land plants.

Liverworts are known for their large chemical diversity. Much of this diversity is synthesized and enclosed within oil bodies (OBs), a synapomorphy of the lineage. OBs contain the enzymes to biosynthesize and store large quantities of sesquiterpenoids and other compounds while limiting their cytotoxicity. Recent important biochemical and molecular discoveries related to OB formation, diversity, and biochemistry allow comparison with other secretory structures of land plants from an evo-devo perspective. This review addresses and discusses the most recent advances in OB origin, development, and function towards understanding the importance of these organelles in liverwort physiology and adaptation to changing environments. Our mapping of OB types and chemical compounds to the current liverwort phylogeny suggests that OBs were present in the most recent common ancestor of liverworts, supporting that OBs evolved as the first secretory structures in land plants. Yet, we require better sampling to define the macroevolutionary pattern governing the ancestral type of OB. We conclude that current efforts to find molecular mechanisms responsible for the morphological and chemical diversity of secretory structures will help understand the evolution of each major group of land plants, and open new avenues in biochemical research on bioactive compounds in bryophytes and vascular plants.

The complete plastid genome sequence of the enigmatic moss, Takakia lepidozioides (Takakiopsida, Bryophyta): evolutionary perspectives on the largest collection of genes in mosses and the intensive RNA editing.

KEY MESSAGE: Complete chloroplast genome sequence of a moss, Takakia lepidozioides (Takakiopsida) is reported. The largest collection of genes in mosses and the intensive RNA editing were discussed from evolutionary perspectives. We assembled the entire plastid genome sequence of Takakia lepidozioides (Takakiopsida), emerging from the first phylogenetic split among extant mosses. The genome sequences were assembled into a circular molecule 149,016 bp in length, with a quadripartite structure comprising a large and a small single-copy region separated by inverted repeats. It contained 88 genes coding for proteins, 32 for tRNA, four for rRNA, two open reading frames, and at least one pseudogene (tufA). This is the largest number of genes of all sequenced plastid genomes in mosses and Takakia is the only moss that retains the seven coding genes ccsA, cysA, cysT, petN rpoA, rps16 and trnPGGG. Parsimonious interpretation of gene loss suggests that the last common ancestor of bryophytes had all seven genes and that mosses lost at least three of them during their diversification. Analyses of the plastid transcriptome identified the extraordinary frequency of RNA editing with more than 1100 sites. We indicated a close correlation between the monoplastidy of vegetative tissue and the intensive RNA editing sites in the plastid genome in land plant lineages. Here, we proposed a hypothesis that the small population size of plastids in each vegetative cell of some early diverging land plants, including Takakia, might cause the frequent fixation of mutations in plastid genome through the intracellular genetic drift and that deleterious mutations might be continuously compensated by RNA editing during or following transcription.

Untargeted In Silico Compound Classification-A Novel Metabolomics Method to Assess the Chemodiversity in Bryophytes.

In plant ecology, biochemical analyses of bryophytes and vascular plants are often conducted on dried herbarium specimen as species typically grow in distant and inaccessible locations. Here, we present an automated in silico compound classification framework to annotate metabolites using an untargeted data independent acquisition (DIA)-LC/MS-QToF-sequential windowed acquisition of all theoretical fragment ion mass spectra (SWATH) ecometabolomics analytical method. We perform a comparative investigation of the chemical diversity at the global level and the composition of metabolite families in ten different species of bryophytes using fresh samples collected on-site and dried specimen stored in a herbarium for half a year. Shannon and Pielou's diversity indices, hierarchical clustering analysis (HCA), sparse partial least squares discriminant analysis (sPLS-DA), distance-based redundancy analysis (dbRDA), ANOVA with post-hoc Tukey honestly significant difference (HSD) test, and the Fisher's exact test were used to determine differences in the richness and composition of metabolite families, with regard to herbarium conditions, ecological characteristics, and species. We functionally annotated metabolite families to biochemical processes related to the structural integrity of membranes and cell walls (proto-lignin, glycerophospholipids, carbohydrates), chemical defense (polyphenols, steroids), reactive oxygen species (ROS) protection (alkaloids, amino acids, flavonoids), nutrition (nitrogen- and phosphate-containing glycerophospholipids), and photosynthesis. Changes in the composition of metabolite families also explained variance related to ecological functioning like physiological adaptations of bryophytes to dry environments (proteins, peptides, flavonoids, terpenes), light availability (flavonoids, terpenes, carbohydrates), temperature (flavonoids), and biotic interactions (steroids, terpenes). The results from this study allow to construct chemical traits that can be attributed to biogeochemistry, habitat conditions, environmental changes and biotic interactions. Our classification framework accelerates the complex annotation process in metabolomics and can be used to simplify biochemical patterns. We show that compound classification is a powerful tool that allows to explore relationships in both molecular biology by "zooming in" and in ecology by "zooming out". The insights revealed by our framework allow to construct new research hypotheses and to enable detailed follow-up studies.

Bryophytes are predicted to lag behind future climate change despite their high dispersal capacities.

The extent to which species can balance out the loss of suitable habitats due to climate warming by shifting their ranges is an area of controversy. Here, we assess whether highly efficient wind-dispersed organisms like bryophytes can keep-up with projected shifts in their areas of suitable climate. Using a hybrid statistical-mechanistic approach accounting for spatial and temporal variations in both climatic and wind conditions, we simulate future migrations across Europe for 40 bryophyte species until 2050. The median ratios between predicted range loss vs expansion by 2050 across species and climate change scenarios range from 1.6 to 3.3 when only shifts in climatic suitability were considered, but increase to 34.7-96.8 when species dispersal abilities are added to our models. This highlights the importance of accounting for dispersal restrictions when projecting future distribution ranges and suggests that even highly dispersive organisms like bryophytes are not equipped to fully track the rates of ongoing climate change in the course of the next decades.

Forest canopy-cover composition and landscape influence on bryophyte communities in Nothofagus forests of southern Patagonia.

Bryophytes (liverworts, mosses and hornworts) are one of the most diverse plant groups worldwide but one of the least studied in temperate forests from an ecological perspective. In comparison to vascular plants, bryophytes have a broader distribution and a longer altitudinal gradient, and their influence on the landscape is poorly understood. The objective was to evaluate environmental drivers that can influence bryophyte cover, richness, diversity, and nestedness in different forest canopy compositions in two typical landscapes across the natural distribution of bryophytes in Tierra del Fuego (Argentina). Three natural Nothofagus forest types (pure deciduous, pure evergreen, and mixed deciduous-evergreen) in two landscapes (coasts < 100 m.a.s.l. and mountains > 400 m.a.s.l.) were selected (N = 60 plots). In each plot, we established one transect (10 m length) to measure bryophyte cover (point-intercept method). Data were evaluated using generalized linear mixed models and multivariate analyses. The studied environmental drivers were mainly explained by the microclimate, with higher effective annual precipitation and relative air humidity in the coastal forests and higher soil moisture in the mountain forests. Greater liverwort richness was found in evergreen forests at the mountain (9 species) than at the coastal, while mosses showed higher richness in mixed deciduous-evergreen forests at the coastal (11 species) than at the mountain. However, the expected richness according to the rarefaction/extrapolation curves suggested that it is possible to record additional species, except for liverworts in pure deciduous forests on the coasts. Similarities and differences among the studied forest types and among plots of the same forest type and landscape were detected. These differences in the studied indexes (similarity that varied between 0 and 1) ranged from 0.09-0.48 for liverworts and 0.05-0.65 for mosses. Moreover, these results indicated that pure evergreen and mixed deciduous-evergreen forests presented higher moss cover (10.7% and 10.0%, respectively), mainly in the mountains than on the coast. These outputs highlight the need to explore differences at greater altitudinal ranges to achieve sustainability objectives conservation planning for bryophytes in southernmost forests.

Optimization of the second internal transcribed spacer (ITS2) for characterizing land plants from soil.

Molecular-based taxonomy, specifically DNA barcoding, has streamlined organism identification. For land plants, the recommended 2-locus barcode of rbcL and matK is not suitable for all groups, thus the second subunit of the nuclear internal transcribed spacer (ITS2) has received attention as a possible alternative. To date, evaluations of ITS2 have mostly been limited in scope to specific plant orders/families and single source material. Prior to using ITS2 to routinely characterize land plants present in environmental samples (i.e., DNA metabarcoding), a wet lab protocol optimized for bulk sample types is needed. To address this gap, in this study we determined the broad recoverability across land plants when using published ITS2 primer pairs, and subsequently optimized the PCR reaction constituents and cycling conditions for the best two performing primer pairs (ITS2F/ITSp4 and ITSp3/ITSu4). Using these conditions, both primer pairs were used to characterize land plants present in 17 diverse soils collected from across the US. The resulting PCR amplicons were prepared into libraries and pooled for sequencing on an Illumina® MiniSeq. Our existing bioinformatics workflow was used to process raw sequencing data and taxonomically assign unique ITS2 plant sequences by comparison to GenBank. Given strict quality criteria were imposed on sequences for inclusion in data analysis, only 43.6% and 7.5% of sequences from ITS2F/ITSp4 and ITSp3/ITSu4 respectively remained for taxonomic comparisons; ~7-11% of sequences originated from fungal co-amplification. The number of orders and families recovered did differ between primer pairs, with ITS2F/ITSp4 consistently outperforming ITSp3/ITSu4 by >15%. Primer pair bias was observed in the recovery of certain taxonomic groups; ITS2F/ITSp4 preferentially recovered flowering plants and grasses, whereas ITSp3/ITSu4 recovered more moss taxa. To maximize data recovery and reduce potential bias, we advocate that studies using ITS2 to characterize land plants from environmental samples such as soil use a multiple primer pair approach.

Reassessing the role of morphology in bryophyte phylogenetics: Combined data improves phylogenetic inference despite character conflict.

Morphological data has gained renewed attention and has been shown to be crucial in clarifying the phylogenetic relationship in a wide range of taxa. In the last decades, phylogenetic analyses of sequence-level data have radically modified the systematic schemes within bryophytes (early non-vascular land plants) and have revealed a widespread pattern of conflict with morphology-based classifications. Yet, a comprehensive evaluation of character conflict has not yet been performed in the context of combined matrices. In this study, we evaluate the impact of morphology on bryophyte phylogeny following a total-evidence approach across 10 published matrices. The analysed matrices span a wide range of bryophytes, taxonomic levels, gene sampling and number of morphological characters and taxa. Data conflict was addressed by measuring: (i) the topological congruence between individual partitions, (ii) changes in support values of the combined data relative to the molecular partition and (iii) clade stability. The association between these measures and the number of morphological characters per taxon (Nc/T ratio) and the proportion of non-fixed characters (i.e., inapplicable, polymorphic and missing data) was explored. In the individual partition analyses, the Nc/T ratio correlated positively with the topological congruence in six to seven datasets depending on the weighting scheme. The proportion of non-fixed cells had a minor influence on congruence between data partitions. The number of characters and proportion of non-fixed data varied significantly between morphological datasets that improved congruence between data types. This variation suggests that morphological datasets affect the results of combined analyses in different ways, depending on the taxa studied. Combined analyses revealed that, despite the low congruence values between partitions, integrating data types improves support values and stability. However, while non-fixed data had no negative effect on support values, stability was reduced as the proportion of non-fixed cells increased. Nc/T ratio was negatively associated with support values and it showed ambiguous responses in stability evaluations. Overall, the results indicate that adding morphology may contribute to the inference of phylogenetic relationships of bryophytes despite character conflict. Our findings suggest that merely comparing (a) morphology-based classifications with molecular phylogenies or (b) the outcome from individual data partitions can misestimate data conflict. These findings imply that analyses of combined data may provide conservative assessments of data conflict and, eventually, lead to an improved sampling of morphological characters in large-scale analyses of bryophytes.

Bryophytes and the symbiotic microorganisms, the pioneers of vegetation restoration in karst rocky desertification areas in southwestern China.

In karst rocky desertification areas, bryophytes coexist with algae, bacteria, and fungi on exposed calcareous rocks to form a bryophyte crust, which plays an irreplaceable role in the restoration of karst degraded ecosystems. We investigated the biodiversity of crust bryophytes in karst rocky desertification areas from Guizhou Province, China. A total of 145 species in 22 families and 56 genera were identified. According to frequency and coverage, seven candidate dominant mosses were screened out, and five drought-resistant indexes of them were measured. Hypnum leptothallum, Racopilum cuspidigerum, and Hyophila involuta have high drought adaptability. We explored the interactions between two dominant mosses (H. leptothallum, H. involuta) and the structure of microbial communities in three karst rocky desertification types. Microbial diversity and function analysis showed that both moss species and karst rocky desertification types affect microbial communities. Moss species much more strongly affected the diversity and changed the community composition of these microbial groups. Bacteria were more sensitive in the microbiome as their communities changed strongly between mosses and drought resistance factors. Moreover, several species of fungi and bacteria could be significantly associated with three drought-resistant indexes: Pro (free proline content), SOD (superoxide dismutase activity), and POD (peroxidase activity), which were closely related to the drought adaptability of mosses. Our results enforced the potential role of moss-associated microbes that are important components involved in the related biological processes when bryophytes adapted to arid habitats, or as one kind of promoters in the distribution pattern of early mosses succession in karst rocky desertification areas.

Mitochondrial genomes of the early land plant lineage liverworts (Marchantiophyta): conserved genome structure, and ongoing low frequency recombination.

BACKGROUND: In contrast to the highly labile mitochondrial (mt) genomes of vascular plants, the architecture and composition of mt genomes within the main lineages of bryophytes appear stable and invariant. The available mt genomes of 18 liverwort accessions representing nine genera and five orders are syntenous except for Gymnomitrion concinnatum whose genome is characterized by two rearrangements. Here, we expanded the number of assembled liverwort mt genomes to 47, broadening the sampling to 31 genera and 10 orders spanning much of the phylogenetic breadth of liverworts to further test whether the evolution of the liverwort mitogenome is overall static. RESULTS: Liverwort mt genomes range in size from 147 Kb in Jungermanniales (clade B) to 185 Kb in Marchantiopsida, mainly due to the size variation of intergenic spacers and number of introns. All newly assembled liverwort mt genomes hold a conserved set of genes, but vary considerably in their intron content. The loss of introns in liverwort mt genomes might be explained by localized retroprocessing events. Liverwort mt genomes are strictly syntenous in genome structure with no structural variant detected in our newly assembled mt genomes. However, by screening the paired-end reads, we do find rare cases of recombination, which means multiple concurrent genome structures may exist in the vegetative tissues of liverworts. Our phylogenetic analyses of the nuclear encoded double stand break repair protein families revealed liverwort-specific subfamilies expansions. CONCLUSIONS: The low repeat recombination level, selection, along with the intensified nuclear surveillance, might together shape the structural evolution of liverwort mt genomes.

Species diversity patterns in managed Scots pine stands in ancient forest sites.

Continuity in forest habitats is crucial for species diversity and richness. Ancient Scots pine forests are usually under forest management, which disturbs vegetation and causes differentiation in terms of tree stand age. To date, vegetation variability in ancient Scots pine forests has not been examined based on tree stand age classes. In the present study the continuity of a large Scots pine forest complex was investigated, and a system of sampling plots established in five tree stand age classes: initiation stands (4-10 years), young stands (20-35 years), middle-aged stands (45-60 years), pre-mature stands (70-85 years) and mature stands (95-110 years). Species composition, including vascular plants, bryophytes and lichens, on soil, tree trunks, and coarse woody debris, was analyzed. Based on existing classifications systems, forest species and ancient forest species groups were distinguished. In the studied ancient Scots pine forests the species pool and richness were relatively low, and the vegetation consisted mostly of generalist species. Cryptogams, which can grow on diverse substrates, were the most abundant species. Moreover, most species could tolerate both forest and non-forest conditions. Age class forests provided different environmental niches for species. Initiation stands were optimal for terrestrial light-demanding species, and in terms of species composition, initiation stands were most specific. Young stands were most preferred by species on coarse woody debris, and at this stage of stand maturation epiphytic species re-appeared. The oldest stands were not rich in forest specialists, i.e. species of closed forest and ancient forest species. Cryptogams of closed forests inhabited different substrates, and they were not associated only with the oldest stands. The low number of forest specialists in the oldest stands may be a general feature of acidophilus pine forests. However, it may also be a result of the lack of species sources in the vicinity of maturing pine stands. In managed forests a frequent diversity pattern is an increase in a species pool and richness after clear-cut logging. In the present study we obtained higher species pools in initiation and young stands, but richness was similar in all tree stand age classes. This resulted from taking into account species of different substrates (terrestrial, epixylous and epiphytic species) which changed their participation in the vegetation of subsequent stages of tree stand development.

Potential of Transcript Editing Across Mitogenomes of Early Land Plants Shows Novel and Familiar Trends.

RNA editing alters the identity of nucleotides in an RNA sequence so that the mature transcript differs from the template defined in the genome. This process has been observed in chloroplasts and mitochondria of both seed and early land plants. However, the frequency of RNA editing in plant mitochondria ranges from zero to thousands of editing sites. To date, analyses of RNA editing in mitochondria of early land plants have been conducted on a small number of genes or mitochondrial genomes of a single species. This study provides an overview of the mitogenomic RNA editing potential of the main lineages of these two groups of early land plants by predicting the RNA editing sites of 33 mitochondrial genes of 37 species of liverworts and mosses. For the purpose of the research, we newly assembled seven mitochondrial genomes of liverworts. The total number of liverwort genera with known complete mitogenome sequences has doubled and, as a result, the available complete mitogenome sequences now span almost all orders of liverworts. The RNA editing site predictions revealed that C-to-U RNA editing in liverworts and mosses is group-specific. This is especially evident in the case of liverwort lineages. The average level of C-to-U RNA editing appears to be over three times higher in liverworts than in mosses, while the C-to-U editing frequency of the majority of genes seems to be consistent for each gene across bryophytes.

Resolution of the ordinal phylogeny of mosses using targeted exons from organellar and nuclear genomes.

Mosses are a highly diverse lineage of land plants, whose diversification, spanning at least 400 million years, remains phylogenetically ambiguous due to the lack of fossils, massive early extinctions, late radiations, limited morphological variation, and conflicting signal among previously used markers. Here, we present phylogenetic reconstructions based on complete organellar exomes and a comparable set of nuclear genes for this major lineage of land plants. Our analysis of 142 species representing 29 of the 30 moss orders reveals that relative average rates of non-synonymous substitutions in nuclear versus plastid genes are much higher in mosses than in seed plants, consistent with the emerging concept of evolutionary dynamism in mosses. Our results highlight the evolutionary significance of taxa with reduced morphologies, shed light on the relative tempo and mechanisms underlying major cladogenic events, and suggest hypotheses for the relationships and delineation of moss orders.

Placing the regionally threatened moss Orthodontium gracile in the big picture - Phylogeny, genome incongruence and anthropogenic dispersal in the order Orthodontiales.

The Orthodontiaceae is a small family of predominantly Southern Hemisphere temperate and South East Asian mosses that has a key phylogenetic position for research into the evolution of pleurocarpy. In the United Kingdom it is represented by the rare conservation priority species Orthodontium gracile and the abundant exotic O. lineare, introduced from the Southern Hemisphere around a century ago. Although the two species are superficially very similar and difficult to tell apart in the field, very little is known about how closely they are related or about the phylogeny, biogeography and evolutionary history of the genus Orthodontium as a whole. Phylogenetic inference and divergence time estimation were used to explore relationships within the genus globally, date major lineage splits, detect reticulate evolutionary processes and test monophyly of taxa. It was shown that Orthodontium gracile belongs to a Holarctic and Asian clade that diverged from the exclusively southern temperate lineage of O. lineare approximately 53 Ma and that it is sister to the Himalayan and South Siberian bispecific genus Orthodontopsis, which we now recognise as a single species within Orthodontium, O. lignicola. Orthodontium lignicola is quite distinct from O. gracile morphologically but may have a closely overlapping centre of extant diversity in the Himalaya, in contrast to O. lineare which is morphologically similar but biogeographically dissimilar. The introduced European populations of Orthodontium lineare were shown to share plastid and nuclear haplotypes with four collections from Tasmania and Southern Chile, but to be distinct from other Chilean and South African haplotypes. Finally, well-supported incongruence between nuclear and plastid sequences in some Western North American populations of Orthodontium gracile strongly implies one or more chloroplast capture or horizontal genome transfer events involving this species and the regionally sympatric O. pellucens. An appeal is made for targeting phylogenetic research at the intersection points of practical conservation, taxonomic uncertainty and wider biological questions and for the factoring of historical evolutionary and phylogenetic diversity into conservation assessments.

Contributions to the bryological knowledge of ASPA 125, Fildes Peninsula, King George Island.

BACKGROUND: With 29 Km2, the Fildes Peninsula is the largest ice free area in King George Island and probably in Antarctica. The region is house of six permanent bases including the only airport in the South Shetlands, which led to impacts on its original landscape and vegetation. In recognition for the need to protect natural values, an Antarctic Specially Protected Area (ASPA 125) was established in the region. Focused mostly on protecting the fossils, the ASPA also plays a role in protecting the vegetation but so far, the management plan for the area does not contain a list of moss species present there. RESULTS: We provided an updated study and checklist of mosses present in ASPA 125. A key to species identification and photographs of main morphological features are also available in this paper. Also, six new occurrences are reported for Fildes Peninsula. CONCLUSION: Considering the scarce knowledge about specific local floras in Antarctica associated with highly impacted area, of which only a fraction is protected, it is suggested the necessity to invest in detailed sampling studies, as well as in a better understanding of the local floras interactions in Antarctica.

Vascular plant and bryophyte species richness in response to water quality in lowland spring niches with different anthropogenic impacts.

Many freshwater ecosystems face severe threats from anthropogenic disturbances. In the field, we investigated the morphology of spring niches and the species richness of vascular plants and bryophytes in 16 springs, draining the quaternary aquifers, located in two different environments-an urban area (city of Bialystok) and a protected area (Knyszyn Forest Landscape Park, NE Poland). In total, 47 vascular plant and 45 bryophyte species were recorded, representing mostly crenophytes including protected species. The most important water quality parameters that can be used to evaluate variations of the spring water chemistry in NE Poland are the mineral-related parameters (electrolytic conductivity, Ca2+, SO42-, and Cl-). The organic-related parameters (DOC) and nutrients (TP, NO3--N) were negatively involved in water quality variations. Our results show that anthropogenic activity significantly affects the biodiversity of plant communities in lowland springs. The presence or absence of crenophytes and bryophytes is indicative of the ecological status of the groundwater outflow complexes.

Gene refashioning through innovative shifting of reading frames in mosses.

Early-diverging land plants such as mosses are known for their outstanding abilities to grow in various terrestrial habitats, incorporating tremendous structural and physiological innovations, as well as many lineage-specific genes. How these genes and functional innovations evolved remains unclear. In this study, we show that a dual-coding gene YAN/AltYAN in the moss Physcomitrella patens evolved from a pre-existing hemerythrin gene. Experimental evidence indicates that YAN/AltYAN is involved in fatty acid and lipid metabolism, as well as oil body and wax formation. Strikingly, both the recently evolved dual-coding YAN/AltYAN and the pre-existing hemerythrin gene might have similar physiological effects on oil body biogenesis and dehydration resistance. These findings bear important implications in understanding the mechanisms of gene origination and the strategies of plants to fine-tune their adaptation to various habitats.

Development of the sphagnoid areolation pattern in leaves of Palaeozoic protosphagnalean mosses.

Background and Aims: Protosphagnalean mosses constitute the largest group of extinct mosses of still uncertain affinity. Having the general morphology of the Bryopsida, some have leaves with an areolation pattern characteristic of modern Sphagna. This study describes the structure and variation of these patterns in protosphagnalean mosses and provides a comparison with those of modern Sphagna. Methods: Preparations of fossil mosses showing preserved leaf cell structure were obtained by dissolving rock, photographed, and the resulting images were transformed to graphical format and analysed with Areoana computer software. Key Results: The sphagnoid areolation pattern is identical in its basic structure for both modern Sphagnum and Palaeozoic protosphagnalean mosses. However, in the former group the pattern develops through unequal oblique cell divisions, while in the latter the same pattern is a result of equal cell divisions taking place in a specific order with subsequent uneven cell growth. The protosphagnalean pathway leads to considerable variability in leaf structure. Conclusions: Protosphagnalean mosses had a unique ability to switch the development of leaf areolation between a pathway unique to Sphagnum and another one common to all other mosses. This developmental polyvariancy hinders attempts to classify these mosses, as characters previously considered to be of generic significance can be shown to co-occur in one individual leaf. New understanding of the ontogeny has allowed us to re-evaluate the systematic significance of such diagnostic characters in these Palaeozoic plants, showing that their similarity to Sphagnum is less substantial.

Biodiversity of biological soil crusts from the Polar Regions revealed by metabarcoding.

Biological soil crusts (BSCs) are amalgamations of autotrophic, heterotrophic and saprotrophic organisms. In the Polar Regions, these unique communities occupy essential ecological functions such as primary production, nitrogen fixation and ecosystem engineering. Here, we present the first molecular survey of BSCs from the Arctic and Antarctica focused on both eukaryotes and prokaryotes as well as passive and active biodiversity. Considering sequence abundance, Bryophyta is among the most abundant taxa in all analyzed BSCs suggesting that they were in a late successional stage. In terms of algal and cyanobacterial biodiversity, the genera Chloromonas, Coccomyxa, Elliptochloris and Nostoc were identified in all samples regardless of origin confirming their ubiquitous distribution. For the first time, we found the chrysophyte Spumella to be common in polar BSCs as it was present in all analyzed samples. Co-occurrence analysis revealed the presence of sulfur metabolizing microbes indicating that BSCs also play an important role for the sulfur cycle. In general, phototrophs were most abundant within the BSCs but there was also a diverse community of heterotrophs and saprotrophs. Our results show that BSCs are unique microecosystems in polar environments with an unexpectedly high biodiversity.